LineageInference

Seurat running and integration

library(Seurat)
library(Matrix)
load("/data/proj/GCB_DvB/humanPublication/pubsplicedfull.Rdata")
load("/data/proj/GCB_DvB/humanPublication/pubunsplicedfull.Rdata")
load("/data/proj/GCB_DvB/humanPublication/networkAnnoFull.Rdata")
metadata <- networkAnnotableFile

Load original clustering

load("/data/proj/GCB_DvB/humanPublication/clusteringmanuscipt.Rdata")
annotationpaper <- networkAnnotableFile
annotationpaper$OriginalClustering <- networkAnnotableFile$seurat_clusters
library(viridis)
library(hues)
sidecols <- iwanthue(length(unique(annotationpaper$OriginalClustering)), 0, 360, 36, 180, 13, 73)
sidecols <- sidecols[as.factor(annotationpaper$OriginalClustering)]
annotationpaper$col <- sidecols
#color matching
colorscheme <- annotationpaper$col
for(i in 1:length(levels(annotationpaper$OriginalClustering))){
  #print(i)
  clsel <- levels(annotationpaper$seurat_clusters)[i]
  colordist <- as.data.frame(table(annotationpaper$col[annotationpaper$seurat_clusters %in% clsel]))
  coloruse <- colordist$Freq
  names(coloruse) <- colordist$Var1
  coloruse <- names(sort(coloruse,decreasing = TRUE))[1]
  colorscheme[which(annotationpaper$seurat_clusters %in% clsel)] <- coloruse
}
annotationpaper$col.translated <- colorscheme
library(plyr)
Level1  <- revalue(as.factor(annotationpaper$OriginalClustering), c(
  "26"="OPC",
  "15"="Glioblast",
  "7"="Glioblast",
  "8"="Glioblast",
  "25"="Glioblast",
  "27"="VLMC",
  "23"="NeuralProgenitor",
  "14"="NeuralProgenitor",
  "6"="NeuralProgenitor",
  "3"="NeuralProgenitor",
  "13"="NeuralProgenitor",
  "17"="NeuralProgenitor",
  "11"="RadialGlia",
  "1"="RadialGlia",
  "4"="RadialGlia",
  "21"="RadialGlia",
  "20"="RadialGlia",
  "18"="RadialGlia",
  "24"="RadialGlia",
  "12"="EarlyInhibitory",
  "5"="EarlyInhibitory",
  "22"="EarlyInhibitory",
  "19"="EarlyInhibitory",
  "2"="EarlyInhibitory",
  "10"="EarlyInhibitory",
  "28"="EarlyInhibitory",
  "9"="EarlyExcitatory",
  "16"="EarlyExcitatory",
  "0"="EarlyExcitatory",
  "29"="Unknown",
  "30"="Unknown",
  "31"="Unknown"
  ))
Level1 <- factor(Level1, levels= c(
  "OPC",
  "Glioblast",
  "RadialGlia",
  "NeuralProgenitor",
  "EarlyInhibitory",
  "EarlyExcitatory",
  "VLMC",
  "Unknown"))
levels(Level1)

[1] "OPC"              "Glioblast"        "RadialGlia"       "NeuralProgenitor" "EarlyInhibitory"  "EarlyExcitatory" 
[7] "VLMC"             "Unknown"         

c("OPC","Glioblast","RadialGlia","NeuralProgenitor","EarlyInhibitory","EarlyExcitatory","VLMC","29","30","31")     

 [1] "OPC"              "Glioblast"        "RadialGlia"       "NeuralProgenitor" "EarlyInhibitory"  "EarlyExcitatory" 
 [7] "VLMC"             "29"               "30"               "31"              

Level1colpal <- c("#E8C688","#D3AC9C","#B9CB6C","#738BC6","#D0D1BD","#DA8B6C","#E0C196","#C9B9A4","#C9B9A4","#C9B9A4") 
Level1col <- Level1colpal[as.factor(Level1)]

annotationpaper$Age <- annotationpaper$Sample
library(plyr)
annotationpaper$Age  <- revalue(as.factor(annotationpaper$Age), c(
  "hF_WK10_2017"="PCW 10",
  "hF_WK10psaNCAM_2017"="PCW 10",
  "hF_WK8_2017"="PCW 8",
  "hF_WK8.5cer.ctx_2019"="PCW 8.5",
  "hF_WK8.5fb_2019"="PCW 8.5",
  "hF_WK9_2019"="PCW 9"))
annotationpaper$Age <- factor(annotationpaper$Age, levels= c(
  "PCW 8",
  "PCW 8.5",
  "PCW 9",
  "PCW 10"))
#merge psaNCAM with non-psaNCAM
annotationpaper$Samples  <- revalue(as.factor(annotationpaper$Sample), c(
   "hF_WK10_2017"="hF_WK10V2",
  "hF_WK10psaNCAM_2017"="hF_WK10V2",
  "hF_WK8_2017"="hF_WK8V2",
  "hF_WK8.5cer.ctx_2019"="hF_WK8.5cer.ctxV3",
  "hF_WK8.5fb_2019"="hF_WK8.5fbV3",
  "hF_WK9_2019"="hF_WK9V3"))

annotationpaper$Level1 <- Level1
annotationpaper$Level1col <- Level1col
colorscheme <- annotationpaper$Level1col
for(i in 1:length(levels(annotationpaper$Level1))){
  #print(i)
  clsel <- levels(annotationpaper$seurat_clusters)[i]
  colordist <- as.data.frame(table(annotationpaper$Level1col[annotationpaper$seurat_clusters %in% clsel]))
  coloruse <- colordist$Freq
  names(coloruse) <- colordist$Var1
  coloruse <- names(sort(coloruse,decreasing = TRUE))[1]
  colorscheme[which(annotationpaper$seurat_clusters %in% clsel)] <- coloruse
}
annotationpaper$Level1col.translated <- colorscheme
colorscheme <- annotationpaper$Level1
for(i in 1:length(levels(annotationpaper$seurat_clusters))){
  #print(i)
  clsel <- levels(annotationpaper$seurat_clusters)[i]
  colordist <- as.data.frame(table(annotationpaper$Level1[annotationpaper$seurat_clusters %in% clsel]))
  coloruse <- colordist$Freq
  names(coloruse) <- colordist$Var1
  coloruse <- names(sort(coloruse,decreasing = TRUE))[1]
  colorscheme[which(annotationpaper$seurat_clusters %in% clsel)] <- coloruse
}
annotationpaper$Level1.translated <- colorscheme

Integrate the version 2 and version 3 data using Seurat CCA

library(Matrix)
# A list of cell cycle markers, from Tirosh et al, 2015, is loaded with Seurat.  We can
# segregate this list into markers of G2/M phase and markers of S phase
s.genes <- cc.genes$s.genes
g2m.genes <- cc.genes$g2m.genes
# Create our Seurat object and complete the initalization steps
hF <- CreateSeuratObject(counts = emats, project = "hFetal", meta.data = metadata)
#split the dataset into a list of two seurat objects
hF.list <- SplitObject(hF, split.by = "version")
hF.list <- lapply(X = hF.list, FUN = function(x) {
    x <- subset(x, subset = nFeature_RNA > 10)
    x <- NormalizeData(x)
    x <- FindVariableFeatures(x, selection.method = "vst", nfeatures = 10000)
})

Performing log-normalization
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[----|----|----|----|----|----|----|----|----|----|
**************************************************|
Calculating gene variances
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**************************************************|
Calculating feature variances of standardized and clipped values
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**************************************************|
Performing log-normalization
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[----|----|----|----|----|----|----|----|----|----|
**************************************************|
Calculating gene variances
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[----|----|----|----|----|----|----|----|----|----|
**************************************************|
Calculating feature variances of standardized and clipped values
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[----|----|----|----|----|----|----|----|----|----|
**************************************************|

#select features that are repeatedly variable across datasets for integration
features <- SelectIntegrationFeatures(object.list = hF.list,nfeatures = 10000)
hF.anchors <- FindIntegrationAnchors(object.list = hF.list, anchor.features = features,reduction = "cca",dims = 1:30)

Scaling features for provided objects

  |                                                  | 0 % ~calculating  
  |+++++++++++++++++++++++++                         | 50% ~05s          
  |++++++++++++++++++++++++++++++++++++++++++++++++++| 100% elapsed=13s  

Finding all pairwise anchors

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Running CCA
Merging objects
Finding neighborhoods
Finding anchors
    Found 23786 anchors
Filtering anchors
    Retained 16240 anchors

  |++++++++++++++++++++++++++++++++++++++++++++++++++| 100% elapsed=03m 12s

#this command creates an 'integrated' data assay
hF <- IntegrateData(anchorset = hF.anchors)

Merging dataset 1 into 2
Extracting anchors for merged samples
Finding integration vectors
Finding integration vector weights
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[----|----|----|----|----|----|----|----|----|----|
**************************************************|
Integrating data

DefaultAssay(hF) <- "integrated"
hF@meta.data$seurat_clusters2 <- hF@meta.data$seurat_clusters
#hF <- NormalizeData(hF)
hF <- FindVariableFeatures(hF, selection.method = "vst",nfeatures = 10000)

Warning in FindVariableFeatures.Assay(object = assay.data, selection.method = selection.method,  :
  selection.method set to 'vst' but count slot is empty; will use data slot instead
Calculating gene variances
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**************************************************|
Warning in eval(predvars, data, env) : NaNs produced
Warning in hvf.info$variance.expected[not.const] <- 10^fit$fitted :
  number of items to replace is not a multiple of replacement length
Calculating feature variances of standardized and clipped values
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**************************************************|

hF <- ScaleData(hF, features = VariableFeatures(hF))

Centering and scaling data matrix

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hF <- RunPCA(hF, features = VariableFeatures(hF), ndims.print = 1:5, nfeatures.print = 3)

PC_ 1 
Positive:  STMN2, MLLT11, RTN1 
Negative:  VIM, HMGB2, ZFP36L1 
PC_ 2 
Positive:  EDNRB, ATP1A2, BCAN 
Negative:  TOP2A, UBE2C, AURKB 
PC_ 3 
Positive:  NFIA, NEUROD6, NFIB 
Negative:  GAD2, PBX3, OTX2 
PC_ 4 
Positive:  RPS6, LINC01551, RPS20 
Negative:  TUBB4B, CCNB1, ARL6IP1 
PC_ 5 
Positive:  OTX2, RPS28, SOX14 
Negative:  MTCO3P12, H3-3A, SP9 

# view cell cycle scores and phase assignments
# Visualize the distribution of cell cycle markers across
hF <- RunUMAP(hF, dims=1:50,n.neighbors = 200,n.components = 2)

23:07:00 UMAP embedding parameters a = 0.9922 b = 1.112
23:07:00 Read 25161 rows and found 50 numeric columns
23:07:00 Using Annoy for neighbor search, n_neighbors = 200
23:07:00 Building Annoy index with metric = cosine, n_trees = 50
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[----|----|----|----|----|----|----|----|----|----|
**************************************************|
23:07:04 Writing NN index file to temp file /tmp/RtmpuZgNaP/file74eb68f0a325
23:07:04 Searching Annoy index using 1 thread, search_k = 20000
23:07:44 Annoy recall = 100%
23:07:46 Commencing smooth kNN distance calibration using 1 thread
23:07:55 Initializing from normalized Laplacian + noise
23:08:07 Commencing optimization for 200 epochs, with 5196278 positive edges
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[----|----|----|----|----|----|----|----|----|----|
**************************************************|
23:09:02 Optimization finished

hF <- CellCycleScoring(hF, s.features = s.genes, g2m.features = g2m.genes, set.ident = TRUE)

Warning: The following features are not present in the object: GINS2, MCM6, MLF1IP, RPA2, UBR7, MSH2, TIPIN, CASP8AP2, CLSPN, POLA1, not searching for symbol synonyms
Warning: The following features are not present in the object: FAM64A, HN1, KIF23, CTCF, CBX5, not searching for symbol synonyms

# hF@meta.data$seurat_clusters2 <- metadata$seurat_clusters
hF <- FindNeighbors(hF, dims = 1:50,k.param = 10)

Computing nearest neighbor graph
Computing SNN

hF <- FindClusters(hF, resolution = 1)

Modularity Optimizer version 1.3.0 by Ludo Waltman and Nees Jan van Eck

Number of nodes: 25161
Number of edges: 425121

Running Louvain algorithm...

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[----|----|----|----|----|----|----|----|----|----|
**************************************************|

Maximum modularity in 10 random starts: 0.9342
Number of communities: 42
Elapsed time: 3 seconds

RidgePlot(hF, features = c("PCNA", "TOP2A", "MCM6", "MKI67"), ncol = 2)

Warning: Could not find MCM6 in the default search locations, found in RNA assay instead
Picking joint bandwidth of 0.128
Picking joint bandwidth of 0.0584
Picking joint bandwidth of 2.99e-06
Picking joint bandwidth of 0.0407

DimPlot(hF,label = TRUE,group.by = "Phase",split.by = "Sample",ncol=2)

DimPlot(hF,label = TRUE,group.by = "Sample",split.by = "version",ncol=2)

DimPlot(hF,label = TRUE,group.by = "version",shuffle = T)

DimPlot(hF,label = TRUE,group.by = "Sample",shuffle = T)

DimPlot(hF,label = TRUE,group.by = "seurat_clusters2")

DimPlot(hF,label = TRUE,group.by = "seurat_clusters")

Write csv to load in python and run scvelo for transition matrix

write.table(hF@assays$integrated@data,file = "/data/proj/GCB_DvB/humanPublication/pubsplicedcorrectedDM.csv",sep=",")
write.table(hF@meta.data,file = "/data/proj/GCB_DvB/humanPublication/pubannoDM.csv",sep=",")
#These are not needed
write.table(hF@reductions$umap@cell.embeddings,file = "/data/proj/GCB_DvB/humanPublication/umapscVeloDM.csv",sep=",")
ematu <- ematu[,colnames(emats)]
# row.names(emats) <- row.names(hF)
# row.names(ematu) <- row.names(hF)
write.table(emats[row.names(hF@assays$integrated@data),colnames(hF)],file = "/data/proj/GCB_DvB/humanPublication/pubsplicedFtruncDM.csv",sep=",")
write.table(ematu[row.names(hF@assays$integrated@data),colnames(hF)],file = "/data/proj/GCB_DvB/humanPublication/pubunsplicedFtruncDM.csv",sep=",")

After running scVELO in python we load the results back in.

library(Matrix)
library(Seurat)
#load("/data/proj/GCB_DvB/humanPublication/Step1vanBruggen5nnCCAFINAL2.Rdata")
load("/data/proj/GCB_DvB/humanPublication/pubsplicedfull.Rdata")
load("/data/proj/GCB_DvB/humanPublication/pubunsplicedfull.Rdata")
emats <- emats#[,dup.cells]
ematu <- ematu#[,dup.cells]
scVtm <- readMM('/data/proj/GCB_DvB/humanPublication/scVELOtransmatFSCCA30NNveloDM.mtx')
lt <- read.csv("/data/proj/GCB_DvB/humanPublication/scVELOLT_FSCCA30NNveloDM.csv")
#Optional read in transcription factor annotations from SCENIC
TFsSCENIC <- read.table(file="/data/proj/GCB_DvB/human/hFetal/motifs-v9-nr.hgnc-m0.001-o0.0.txt",sep="\t",stringsAsFactors = F)
TFs <- unique(TFsSCENIC$V6)

Now we create the diffusion map integrating both velocity and transcriptional distances into one manifold

set.seed(1234)
#First we create the diffusion map based on transcription
tsnerlba <- t(hF@reductions$pca@cell.embeddings[,1:50])
#get destiny, removed from bioconductor (not very useful >:())
# install.packages("remotes")
# remotes::install_github("https://github.com/theislab/destiny",build_vignettes = FALSE, dependencies = TRUE)
#check for duplicated cells 
# dup.cells <- !duplicated(t(tsnerlba))
# table(dup.cells)
# dup.cells <- as.logical(dup.cells)
# hF <- hF[,dup.cells]
# tsnerlba <- tsnerlba[,dup.cells]

library(destiny)
tsmanifolddist <- DiffusionMap(data =t(tsnerlba) ,n_eigs =20,verbose = TRUE)#,sigma=0.001) 

finding knns......done. Time: 186.81s
Calculating transition probabilities......done. Time: 0.38s

performing eigen decomposition......done. Time: 5.69s

colnames(tsmanifolddist@transitions) <- colnames(hF)
row.names(tsmanifolddist@transitions) <- colnames(hF)

#Now we create the joint representation
XY <- t(scVtm) %*% tsmanifolddist@transitions

#Now we integrate the spaces
library(irlba)
dcompl <- irlba(XY,50,maxit = 100000)
tsnerlba <- t(dcompl$u)
tsnerlba <- rbind(tsnerlba,t(dcompl$v))
library(destiny)
tsmanifold <- DiffusionMap(data =t(tsnerlba) ,n_eigs =20,verbose = TRUE)#,sigma=0.001) 

finding knns......done. Time: 149.84s
Calculating transition probabilities......done. Time: 0.31s

performing eigen decomposition......done. Time: 21.49s

colnames(tsmanifold@transitions) <- colnames(hF)
row.names(tsmanifold@transitions) <- colnames(hF)

#Now we generate the backward kernel 80% velocity 20% transcriptional distance
scVtmb <-  scVtm
scVtmb <- t(t(scVtmb)/Matrix::colSums(scVtmb,na.rm = TRUE))
scVtmb[is.na(scVtmb)] <- 0
kernel <- t((scVtm *0.8)) + (tsmanifold@transitions * 0.2)
kernel <- t(t(kernel)/Matrix::colSums(kernel,na.rm = TRUE))
kernel[is.na(kernel)] <- 0

Now we calculate the new clusters based on the new manifold

newclustobj <- Seurat::CreateSeuratObject(counts = as.matrix(hF@assays$RNA@data))
NetworkDist <- as.dist(1-tsmanifold@transitions)
newclustobj[["HSphere_snn"]] <- Seurat::FindNeighbors(NetworkDist,k.param=10)$snn

Building SNN based on a provided distance matrix
Computing SNN
Warning: Adding a Graph without an assay associated with it

newclustobj <- Seurat::FindClusters(newclustobj, graph.name = "HSphere_snn",resolution = 1.6) #algorithm = 4,

Modularity Optimizer version 1.3.0 by Ludo Waltman and Nees Jan van Eck

Number of nodes: 25161
Number of edges: 251057

Running Louvain algorithm...

0%   10   20   30   40   50   60   70   80   90   100%
[----|----|----|----|----|----|----|----|----|----|
**************************************************|

Maximum modularity in 10 random starts: 0.9369
Number of communities: 78
Elapsed time: 1 seconds

Warning: Adding a command log without an assay associated with it

table(newclustobj@meta.data$seurat_clusters)

  0   1   2   3   4   5   6   7   8   9  10  11  12  13  14  15  16  17  18  19  20  21  22  23  24  25  26  27  28  29 
899 873 696 666 570 558 543 531 524 523 516 506 504 498 493 491 478 465 459 442 439 417 414 403 398 387 375 372 368 361 
 30  31  32  33  34  35  36  37  38  39  40  41  42  43  44  45  46  47  48  49  50  51  52  53  54  55  56  57  58  59 
357 355 348 344 343 342 337 337 334 317 317 308 291 290 288 281 281 278 258 257 251 235 232 230 222 191 178 176 169 169 
 60  61  62  63  64  65  66  67  68  69  70  71  72  73  74  75  76  77 
159 154 147 145 138 122 116 115  84  80  74  74  72  65  58  34  29  10 

hF@meta.data$seurat_clusters3 <- newclustobj@meta.data$seurat_clusters
DimPlot(hF,label = TRUE,group.by = "seurat_clusters3")+ NoLegend()

Idents(hF) <- 'seurat_clusters3'
  DimPlot(hF, label=F, cells.highlight=WhichCells(hF, idents = "69"), cols.highlight = c("darkblue", "darkred"), cols= "grey")

  FeaturePlot(hF,features = c("APOD"),slot="data")

And we calculate the pseudotimes associated to each cluster

pt <- destiny::DPT(tsmanifold)#,tips = c(1,2))
cd_diffusionplot <- hF@meta.data
lineagecluster <- as.factor(hF$seurat_clusters3)
names(lineagecluster) <- row.names(cd_diffusionplot)
x<- pt$DPT1
names(x) <- row.names(cd_diffusionplot)
branchtips = as.character(NULL)#lowestcell
branchpseudotimes <- matrix(nrow=(length(levels(as.factor(lineagecluster)))),ncol=nrow(cd_diffusionplot))
lineagecluster <- lineagecluster[row.names(cd_diffusionplot)]
for(i in 1:length(levels(droplevels(as.factor(lineagecluster))))){
  pseudotime <- x[names(lineagecluster[lineagecluster %in% levels(lineagecluster)[i]])]
  topcell <- names(pseudotime[pseudotime == max(pseudotime)])
  branchtips <-  c(branchtips,topcell)
x <-pt[,which(row.names(cd_diffusionplot)==topcell,arr.ind = TRUE)]
names(x) <- row.names(cd_diffusionplot)
  branchpseudotimes[i,] <- x
  #print(i)
}
colnames(branchpseudotimes) <- row.names(cd_diffusionplot)
row.names(branchpseudotimes) <- levels(as.factor(lineagecluster))
branchpseudotimes <- branchpseudotimes[as.character(sort(as.integer(row.names(branchpseudotimes)),decreasing = F)),]

x <- hF@reductions$umap@cell.embeddings[,1]
  y <- hF@reductions$umap@cell.embeddings[,2]
  
load("/data/proj/GCB_DvB/humanPublication/umapharmony.Rdata")
  x <- umap_out_harmony2ddist$layout[,1]
  y <- umap_out_harmony2ddist$layout[,2]

Now we add tracks of lineage diffusion giving the start states of the diffusion process #this is the one to use

#library(e1071)
library(Matrix)
library(destiny)
startpt <- lt$latent_time
names(startpt) <- colnames(branchpseudotimes)
traceclusters <- matrix(ncol=nrow(branchpseudotimes),nrow=ncol(branchpseudotimes))
for(i in 1:nrow(branchpseudotimes)){
  #print(i)
log<- numeric()
starttime = "-1"
if(!starttime == row.names(branchpseudotimes)[i]){
source <- startpt
sink <- branchpseudotimes[i,]
sink <- sort(sink)
trackcellstart <- names(head(sink,10))
lineagecells <- trackcellstart
trackcell <- trackcellstart[1]
threshold <- source[trackcellstart[1]]
stall = 1
while(source[trackcell] > 0 & stall > 0 ) {
  logold <- log
  log<- c(log,source[trackcellstart])
  stall = length(log) - length(logold)
pool <- names(head(sink[sink > sink[trackcell]],1))
if(length(pool) < 1) {trackcell <- names(which(source == min(source)))}
else{
trackcellstart <- names(which(source[pool] <= min(log)))
lineagecells <- c(lineagecells,trackcellstart)
trackcell <- names(which(source[pool] == min(source[pool])))}
}
LineageCells <- lineagecells
Pathtrace <- names(lineagecluster) %in% c(unique(LineageCells))#,unique(LineageCells2))
}
else{Pathtrace <- logical(ncol(branchpseudotimes)) }
traceclusters[,i] <- Pathtrace
}

Here we start the diffusion process, first add the function

WAdiffuse <- function(A,dist,ncells,pool=FALSE,cores,distselectneighbor,nameselectneighbor){
  library(Matrix)
  library(foreach)
  library(doSNOW)
  library(matrixStats)
  range01 <- function(x){(x-min(x))/(max(x)-min(x))}
 cl <- makeCluster(cores)  
 registerDoSNOW(cl)  
iterations <- ncol(A)
pb <- txtProgressBar(max = iterations, style = 3)
progress <- function(n) setTxtProgressBar(pb, n)
opts <- list(progress = progress)
#distselectneighbor <- Matrix(apply(dist,2,function(x) sort(x,decreasing=FALSE)[1:ncells]),sparse = TRUE)
#nameselectneighbor <- apply(dist,2,function(x) sort(x,decreasing=FALSE,index.return=TRUE)$ix[1:ncells])
rm(dist)
gc()
if(class(A) != "dgCMatrix"){
A <- Matrix(as.matrix(A),sparse = T)
}
gc()
if(pool==FALSE){
D <-  foreach(i = 1:ncol(A),.combine=cbind,.packages=c("Matrix"),.options.snow = opts) %dopar% {
distsub <- 1-distselectneighbor[,i] #uses prob
C <- A[,nameselectneighbor[,i]]
C <- apply(C,1,function(x) weighted.mean(x,distsub))
return(C)
}
close(pb)
 stopCluster(cl)
}
if(pool==TRUE){
D <-  foreach(i = 1:ncol(A),.combine=cbind,.packages=c("Matrix"),.options.snow = opts) %dopar% {
distsub <- rowSums(A[,nameselectneighbor[,i]])
return(distsub)
}
close(pb)
 stopCluster(cl)
}
colnames(D) <- colnames(A)
row.names(D) <- row.names(A)
D <- Matrix(D,sparse=TRUE)
  gc()
 return(D)
}

Now we run the diffusion for 30 iterations with 100 neighbors, and calculate lineage membership and diffusion score

library(Matrix)
nameselectneighbor <- apply(kernel,2,function(x) sort(x,decreasing=TRUE,index.return=TRUE)$ix[1:100])
distselectneighbor <- Matrix(apply(kernel,2,function(x) sort(x,decreasing=TRUE)[1:100]),sparse = TRUE)

linwalk <- list()
linwalk[[1]] <- WAdiffuse(t(1*traceclusters),kernel,pool = FALSE,ncells=100,cores=8,distselectneighbor,nameselectneighbor)

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Warning in if (class(A) != "dgCMatrix") { :
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for(i in 2:30){
  print(i)
linwalk[[i]] <- WAdiffuse(linwalk[[i-1]],kernel,pool = FALSE,ncells=100,cores=8,distselectneighbor,nameselectneighbor)
}

[1] 2

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#save(linwalk,file="/data/proj/GCB_DvB/humanPublication/LinwalkFTRUECCAdiff30NNusingkernel.Rdata")
#load("/data/proj/GCB_DvB/humanPublication/LinwalkFTRUECCAdiff30NNusingkernel.Rdata")

#get lineages
for(j in 1:nrow(linwalk[[1]])){
  fpass <- linwalk[[1]][j,]
for(i in 2:30){
fpass <- cbind(fpass,linwalk[[i]][j,])
}
values <- apply(fpass,1,max)
if(j == 1){lineage = values} else{lineage <- rbind(lineage,values)}
}

Get lineage KL function

KLlin <- function(A,B,cores,skipgenes,limitcells,ncells,correct=FALSE,batch){
  print(correct)
  library(Matrix)
  library(foreach)
  library(doSNOW)
  range01 <- function(x){(x-min(x))/(max(x)-min(x))}
 cl <- makeCluster(cores)  
 registerDoSNOW(cl)  
iterations <- ncol(A)
pb <- txtProgressBar(max = iterations, style = 3)
progress <- function(n) setTxtProgressBar(pb, n)
opts <- list(progress = progress)
if(class(A) != "dgCMatrix"){
A <- Matrix(as.matrix(A),sparse = T)
 }
 if(class(B) != "dgCMatrix"){
B <- Matrix(as.matrix(A),sparse = T)
 }
if(min(A) < 0){
A <- A-min(A)
}
if(min(B) < 0){
B <- B-min(B)
}
rm(B)
gc()
D <-  foreach(i = 1:ncol(A),.combine=cbind,.packages="Matrix",.options.snow = opts) %dopar% {
D <- rep(NA, ncol(A))
if(skipgenes==TRUE){
avoidgene <- which(! (A[,i]<=0));C <- A[avoidgene,]}else{C <- A}
Ckl   <- t(t(C)/colSums(C))#[avoidgene,]
Ckl[is.na(Ckl)] <- 0
Ckl <- Matrix(Ckl+0.0000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000001,sparse=TRUE)
Ckl_log <- log(Ckl[,i]/Ckl[,-i])
C1kl <- Ckl[,i]*Ckl_log
rm(Ckl_log)
Ckl <- colSums(C1kl)

nnsort <- sort(Ckl,decreasing = FALSE)

if(limitcells==TRUE){
G <- t(C1kl[,names(nnsort)[1:ncells]]) / Ckl[names(nnsort)[1:ncells]];G <- (t(G) %*% Ckl[names(nnsort)[1:ncells]])[,1]}else{G <- t(C1kl) / Ckl;G <- (t(G) %*% Ckl)[,1]}

GD <- rep(NA, nrow(A))
usedG <- which(row.names(A) %in% names(G))
notusedG <- which(!row.names(A) %in% names(G))
GD[usedG] <- G
GD[notusedG] <- 0
names(GD) <- row.names(A)

C <- GD

gc()
return(C)
}
close(pb)
 stopCluster(cl)
 
colnames(D) <- colnames(A)
rownames(D) <- row.names(A)
Dnorm <- scale(abs(D), center=FALSE, scale=colSums(abs(D)))
Dnorm.sign <- scale(D, center=FALSE, scale=colSums(abs(D)))
D <- Matrix(D,sparse=TRUE)
Dnorm <- Matrix(Dnorm,sparse=TRUE)
Dnorm.sign <- Matrix(Dnorm.sign,sparse=TRUE)
D <- list(D,Dnorm,Dnorm.sign)
names(D) <- c("D","Dnorm","Dnorm.sign")
  #D <- D/sum(D)
  gc()
 return(D)
}

Run lineage membership estimation for each cell in the matrix

lineageKL <- lineage
lineageKLdone <- KLlin(lineage,lineage,15,skipgenes = FALSE,limitcells = FALSE,ncells = 100,correct = FALSE,batch = colnames(hF))

[1] FALSE

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Warning in if (class(B) != "dgCMatrix") { :
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#save(lineageKL,file="/data/proj/GCB_DvB/humanPublication/Final/lineagKLfinalrealCCA30NNvelokernel.Rdata")
#load("/data/proj/GCB_DvB/humanPublication/Final/lineagKLfinalrealCCA30NNvelokernel.Rdata")
lineage.mem <- lineageKLdone$Dnorm
lineage.mem.max <- (apply(lineage.mem,2,max))

Visualize lineage information

library(plotly)
library(dplyr)
#Differentiation score
plot_ly( x = ~x, y = ~y,marker = list(size = 5)) %>% add_markers(color=~lineage.mem.max,alpha=1)

#Plot OL lineage cluster 69 , row 70.
Expression_capped <- lineage.mem[70,]^0.1
Expression_capped <- scale(Expression_capped)
Expression_capped[Expression_capped <= 1.96]  <- 0
    names(Expression_capped) <- seq_len(length(Expression_capped))
    Expression_capped <- sort(Expression_capped,decreasing = FALSE)
    ord <- as.integer(names(Expression_capped))
    plot_ly( x = ~x[ord], y = ~y[ord],marker = list(size = 5)) %>% add_markers(color=~Expression_capped,alpha=1)%>% layout(
      xaxis = list( showgrid = FALSE,showaxis=FALSE,zeroline=FALSE),
      yaxis = list(showgrid = FALSE,showaxis=FALSE,zeroline=FALSE))

library(Matrix)
emat.norm <- hF@assays$RNA@data
#REMEMBER its PLUS1 for the clusters
lnum <- as.numeric(70)
Expression_capped <- lineage.mem[lnum,]^0.1
Expression_capped <- scale(Expression_capped)
#1.645 10% 1.96 5% 2.58 1%  
Expression_capped[Expression_capped <= 1.96]  <- 0
ptlin <- which(Expression_capped > 0)
length(ptlin)

[1] 1051

Lineagegenes <- row.names(hF@assays$integrated@data)
#Only TFs
#Lineagegenes <- Lineagegenes[Lineagegenes %in% TFs]
suppressWarnings(corlin <- apply(as.matrix(emat.norm[Lineagegenes,ptlin]),1,function(x) cor.test(x,lineage.mem[lnum,ptlin]^0.1,method="spearman")))
linKLdrivers <- as.data.frame(t(as.data.frame(lapply(corlin,unlist))))
linKLdrivers$p.value <- as.numeric(as.character(linKLdrivers$p.value))
linKLdrivers$estimate.rho <- as.numeric(as.character(linKLdrivers$estimate.rho))
#linKLdrivers$p.adjust <- p.adjust(linKLdrivers$p.value,method = "BH")
ldsig <- linKLdrivers$p.value;ldmaxcor <-linKLdrivers$estimate.rho
names(ldmaxcor) <- Lineagegenes
names(ldsig) <- Lineagegenes
lineagevals <- as.data.frame(cbind(ldsig,ldmaxcor))
lineagevals <- subset(lineagevals,lineagevals$ldsig < 0.05)
lvgenes <- lineagevals
Lineagegenes <- row.names(lineagevals[order(abs(lineagevals$ldmaxcor),decreasing = TRUE),])#[1:500]


linclusters <- as.data.frame(table(hF$seurat_clusters3[ptlin])/table(hF$seurat_clusters3))
linclusters <- as.character(linclusters[linclusters$Freq >0.1,]$Var1)
corclusters <- hF$seurat_clusters3#[ptlin]
 for(i in 1:length(linclusters)){
  if(i==1){cat("Correlating genes...binning by clusters...")}
  cat(paste("Bin Cluster ",linclusters[i],"..."))
  if(i!=1){linKLdriverssig <-cbind(linKLdriverssig,linKLdrivers$p.value);linKLdriverscor <-cbind(linKLdriverscor,linKLdrivers$estimate.cor)}
binlin  <- emat.norm[Lineagegenes,]
binlin  <- binlin[,corclusters %in% linclusters[i]]
cat(paste(length(which(corclusters %in% linclusters[i])),"cells..."))
  linbin <- lineage.mem[lnum,]^0.1
  linbin <- linbin[corclusters %in% linclusters[i]]
suppressWarnings(corlin <- apply(as.matrix(binlin),1,function(x) cor.test(x,linbin,method="pearson")))
linKLdrivers <- as.data.frame(t(as.data.frame(lapply(corlin,unlist))))
linKLdrivers$p.value <- as.numeric(as.character(linKLdrivers$p.value))
linKLdrivers$p.value <- p.adjust(linKLdrivers$p.value,method = "BH")
linKLdrivers$estimate.cor <- as.numeric(as.character(linKLdrivers$estimate.cor))
if(i!=1){linKLdriverssig <-cbind(linKLdriverssig,linKLdrivers$p.value)}
if(i==1){linKLdriverssig <- linKLdrivers$p.value;linKLdriverscor <-linKLdrivers$estimate.cor}
 }

Correlating genes...binning by clusters...Bin Cluster  6 ...543 cells...Bin Cluster  16 ...478 cells...Bin Cluster  24 ...398 cells...Bin Cluster  39 ...317 cells...Bin Cluster  49 ...257 cells...Bin Cluster  69 ...80 cells...Bin Cluster  73 ...65 cells...

row.names(linKLdriverscor) <- Lineagegenes
row.names(linKLdriverssig) <- Lineagegenes
ldmaxcor <- apply(linKLdriverscor,1,max,na.rm = TRUE)
ldmaxvar <- apply(linKLdriverscor,1,var,na.rm = TRUE)
names(ldmaxcor) <- Lineagegenes
names(ldmaxvar) <- Lineagegenes
ldsig <- apply(linKLdriverssig,1,min,na.rm = TRUE)
names(ldsig) <- Lineagegenes
lineagevals <- as.data.frame(cbind(ldsig,ldmaxcor,ldmaxvar))
lineagevals <- subset(lineagevals,lineagevals$ldsig < 0.05)
lvgenes <- lineagevals
EC <- lineage[lnum,ptlin]
names(EC) <- seq_len(length(EC))
EC <- sort(EC,decreasing = FALSE)
ord <- as.integer(names(EC))

if(nrow(lineagevals) <=100){ptgenes <- row.names(lineagevals)}else{ptgenes <- row.names(lineagevals[order(lineagevals$ldmaxcor,decreasing = TRUE),])[1:100]}

ctlin <- as.matrix(emat.norm[ptgenes,ptlin])
dat <- data.frame(t(ctlin[,ord]))
## response
vars <- colnames(dat)
## covariate
id <- 1:nrow(dat)
library(mgcv)
dat <- data.frame(t(ctlin[,ord]))
colnames(dat) <- gsub("-", "", colnames(dat))
ptgenesnum <- colnames(dat)
dat <- cbind(id,dat)

new.datp <- matrix(nrow=length(ptgenesnum),ncol=length(id))
new.datpval <- numeric(length=length(ptgenesnum))
for(i in 1:length(ptgenesnum)){
  form <- as.formula(paste0(ptgenesnum[i],"~",paste0("s(","id",',bs="cs",k=10',")",collapse="+")))
  m <- gam(form, data=dat)
  form <- as.formula(paste0(ptgenesnum[i],"~",paste0("1",collapse="+")))
  m0 <- gam(form, data=dat)
  new.datp[i,] <- m$fitted.values
}
row.names(new.datp) <- ptgenes
colnames(new.datp) <- colnames(ctlin[,ord])
new.datp[new.datp<0] <- 0
range01 <- function(x){(x-min(x))/(max(x)-min(x))}
new.datp <- t(apply(new.datp,1,range01))
new.dat <- new.datp#[fdr < 0.05,]

GC <- unlist(apply(new.dat,1,function(x) which(x == max(x))[1]))
names(GC) <- seq_len(length(GC))
GC <- sort(GC,decreasing = TRUE)
ordG <- as.integer(names(GC))

ctlin <- as.matrix(ematu[row.names(new.dat),ptlin])
dat <- data.frame(t(ctlin[,ord]))
vars <- colnames(dat)
id <- 1:nrow(dat)
library(mgcv)
dat <- data.frame(t(ctlin[,ord]))
colnames(dat) <- gsub("-", "", colnames(dat))
ptgenesnum <- colnames(dat)
dat <- cbind(id,dat)
new.datu <- matrix(nrow=length(ptgenesnum),ncol=length(id))
for(i in 1:length(ptgenesnum)){
  #print(i)
  form <- as.formula(paste0(ptgenesnum[i],"~",paste0("s(","id",',bs="cs",k=10',")",collapse="+")))
  new.datu[i,] <- gam(form, data=dat)$fitted.values
}
row.names(new.datu) <- row.names(new.dat)
colnames(new.datu) <- colnames(ctlin[,ord])
new.datu[new.datu<0] <- 0
range01 <- function(x){(x-min(x))/(max(x)-min(x))}
new.datu <- t(apply(new.datu,1,range01))
new.dat[new.dat<0] <- 0
new.datu <- new.dat[ordG,]-new.datu[ordG,]
filter <- apply(new.datu,1,anyNA)
new.datu <- new.datu[!filter,]
ordF <- ordG[!filter]
GC <- unlist(apply(new.datu,1,function(x) which(x == max(x))[1]))
names(GC) <- seq_len(length(GC))
GC <- sort(GC,decreasing = TRUE)
ordF <- as.integer(names(GC))


library(heatmap3)
library(pheatmap)
library(hues)
library(viridis)
sidecols <- iwanthue(length(unique(hF@meta.data$seurat_clusters3)),0, 360, 36, 180, 13, 73)
sidecols <- sidecols[as.factor(hF@meta.data$seurat_clusters3)][ptlin]
names(sidecols) <- seq_len(length(sidecols))
sidecols <- sidecols[ord]
heatmap3(new.dat[ordG,], Rowv = NA, Colv = NA ,scale = "none",symm = F, method = "ward.D2",col=viridis(1024),balanceColor =F,cexRow = 1.2,cexCol = 0.5,ColSideColors=sidecols,labCol = NA,)

heatmap3(new.datu[ordF,], Rowv = NA, Colv = NA ,scale = "none",symm = F, method = "ward.D2",balanceColor =F,cexRow = 1.4,cexCol = 0.5,ColSideColors=sidecols,labCol = NA)

save.image("/data/proj/GCB_DvB/humanPublication/Final/NBpart2.RData")